A New Species of Zamia from Chocó, Colombia (Cycadales, Zamiaceae)
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A new species of Zamia from Chocó, Colombia (Cycadales, Zamiaceae) 1 2 1 MICHAEL CALONJE ,DENNIS STEVENSON ,CLAUDIA CALONJE , 3 2 YAN ARLEY RAMOS , AND ANDERS LINDSTROM 1 Author for correspondence: Montgomery Botanical Center, 11901 Old Cutler Road, Miami, FL 33156, U.S.A.; e-mail: [email protected] 2 The New York Botanical Garden, Bronx, NY 10458-5126, U.S.A. 3 Universidad Tecnológica del Chocó, Herbario CHOCO, Grupo de Investigación de la Flora Chocoana, Quibdó, Chocó, Colombia. Abstract. Zamia pyrophylla, a new species from Chocó, Colombia, is described and illustrated. It is compared to Z. amazonum, Z. cunaria, and Z. ipetiensis.Itisdist- inguished by its bright orange and red emerging leaves with leaflets that progress- ively turn green from the apex to the base as they mature, petioles with branched prickles and with ferruginous pubescence when immature, villous strobilar axes, and adaxial microsporangia. Key Words: cycad, Chocó, Zamia. Stevenson (2001) recognized 18 species of A research expedition led by the CHOCO cycads in Colombia, 16 of these included in the herbarium of the Universidad Tecnológica del genus Zamia L., and two in the endemic genus ChocóinQuibdóincollaborationwithMont- Chigua D. W. Stev. Calderón-Sáenz and Steven- gomery Botanical Center (Miami, Florida) took son (2003) subsequently described Zamia oli- place in March of 2009 with the purpose of godonta, bringing the total of recognized species gaining a better understanding of this relatively in Colombia to 19. Six of these species (Z. unknown Zamia from the Chocó. Close exami- amplifolia W. Bull ex. Mast., Z. chigua Seem., nation of several hundred plants across five Z. disodon D.W. Stev. & Sabato, Z. obliqua A. different populations, including fertile plants Braun, and Z. roezlii Linden) occur in the Chocó with ovulate and pollen strobili, led us to the biogeographic region of Colombia. This region conclusion that this taxon represents a new stretches across the entire Pacific coast of species, formally described below. Colombia, including parts of the Departments of Chocó, Valle del Cauca, Cauca, Risaralda, Zamia pyrophylla Calonje, D.W. Stev. & A. Antioquia, Córdoba, and Nariño and is consid- Lindstr. sp. nov. Type: Colombia. Chocó: ered one of the most biologically diverse areas Mun. Quibdó, 60–100m, Calonje et al. COL09– on the planet. In addition to the species noted 014 (holotype: CHOCO; isotypes: COL, above, Stevenson (2001) also listed three Zamia CHOCO, CUVC, HUA). (Figs. 1, 2) collections made in Chocó (Bernal et al. 1090, COL; Juncosa 1721, JAUM, MO, NY; Santa et Zamia cunaria D. W. Stev. simulans, sed marginibus al. 1149, HUA.) under specimens examined for foliolii integerrimis et apice megasporophylli sterilis Zamia amazonum D. W. Stev., noting that the longate differt. material differed somewhat from the typical material and suggesting that further research Stem hypogeous, rarely branching, globose would be necessary to confirm whether this to cylindrical, to 10 cm or more in diameter. material represented a different species. Cataphylls chartaceous, triangular to narrowly Brittonia, 62(1), 2010, pp. 80–85 ISSUED: 1 March 2010 © 2009, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. 2010] CALONJE ET AL.: ZAMIA (ZAMIACEAE) 81 FIG.1. Vegetative features of Zamia pyrophylla. A. Seedling with eophyll and apogeotropic roots. B. Side view of characteristic leaf. C. Frontal view of emergent leaf. D. Cataphylls. E. Median leaflet. F. Caudex of juvenile plant. All photographs taken of plants from type locality (Calonje et al. COL09–014). 82 BRITTONIA [VOL 62 FIG.2. Reproductive features of Zamia pyrophylla. A. Ovulate strobilus at near receptivity stage. B. Mature ovulate strobilus. C. Mature megasporophyll face, frontal view. D. Mature megasporophyll, abaxial view. E. Mature seed with sarcotesta. F. Sclerotesta of mature seed. G. Pollen strobilus at pollen shedding. H. Mature pollen strobilus cross-section, adaxial view. I. Mature pollen strobilus cross-section, abaxial view. J. Juvenile pollen strobili. K. Microsporophyll, adaxial view. L. Microsporophyll, abaxial view. All photographs taken of plants from type locality (Calonje et al. COL09–014). 2010] CALONJE ET AL.: ZAMIA (ZAMIACEAE) 83 triangular, to 8.5×3.0 cm, tan-tomentose with and white hairs, sterile apex 19–20 mm deep brown papyraceous wings. Ptyxis inflexed. with hexagonal to oblong-hexagonal distal face Leaves 1–2, 1.1–3.4 m long; petioles 12.5–13.2 x 15.5–16.2 mm, extruded to a 0.75–2.4 m long with abruptly swollen base to small, depressed terminal facet 2.5–3.4× 4 cm wide, densely covered with stout, some- 5.7–6.3 mm, megasporophyll face tomentose, times branching prickles to 5 mm long; rachis burnt amber in new strobili, rust colored when 0.3–1 m long, curved outward, with prickles in near receptivity, at maturity black with some lower half, petiole and rachis brown tomentose exposed areas glabrous green. Seeds ovoid, when emerging, gradually becoming glabrous. 10–15.6 mm×9.5–13.2 mm, sarcotesta red and Leaflets 8–16 pairs, chartaceous to coriaceous, fleshy, eventually becoming evenly papyra- median ones 2–5.8 cm apart, insertion on rachis ceous at maturity. 6–10 mm wide, lanceolate to oblanceolate and Distribution and habitat.—This species is straight to slightly falcate, alternate to subop- known only from the Atrato Valley in the posite, apex acute to acuminate, margins entire, Municipalities of Quibdó and Atrato, Chocó. basal leaflets 28–51×2.2–7.2 cm; median leaf- It occurs in tropical rainforest at elevations lets 24.5–55×4–7.1 cm; apical leaflets between 10–100 m. Because it has a sub- 19–34.5×2.1–6.6 cm wide, maroon at emer- terranean stem, it is able to survive initial gence, turning to orange or reddish tones, and forest clearing and can thrive and produce gradually turning green from the leaflet apex to strobili while growing in full sun in pastures the base as they mature. Eophylls with unarmed if left undisturbed. Plants were frequently petioles 19–25 cm long, rachis to 1 cm long, observed with emergent new leaves in typically carrying four leaflets 6.1–6.5× recently cleared fields, but were absent from 1.7–2.2 cm. Pollen strobili conical-cylindrical, older clearings, suggesting the plants may not erect or slightly spreading, 5–15 +, at pollen be able to survive repeated leaf excision from shedding 7.1–8.1×1.5–1.7 cm, strobilar axis pasture clearing. and proximal section of microsporophyll vil- The area near Quibdó is one of the wettest lous with mixed white and rust-brown hairs, on earth, with an average annual precipitation peduncles tomentose, emerging white to cream of 8558 mm (West, 1957). The drier season and maturing maroon to brown, 14–25× occurs from January through March with 0.8–1 cm, microsporophylls spirally arranged February being the driest month, and the in 8–10 orthostichies of 13–20 sporophylls rainy season occurs April through December, each, obtrullate, 5.9–6.3×3.5–4.3 mm at pollen with August being the wettest month. The shedding, sterile apex encompassing 1/4 to 1/2 temperature averages 28°C and ranges from of total length of a microsporophyll, tomentose, 15°C to 41°C (Forero, 1982). tan to cream colored in proximal half, speckled Reproductive phenology.— Very little is maroon to brown in distal half, face hexagonal known about the reproductive phenology of to oblong hexagonal, 2.7–3×3.5–4 mm, this species. All herbarium specimens exam- extruded downward to a narrow horizontal ined prior to our March 2009 expedition were facet, abaxial side of microsporophyll with sterile with no mention of reproductive 15–20 microsporangia limited to the proximal structures in the label notes. During this half and along the margins, adaxial side with expedition, we were fortunate to observe both 12–15 microsporangia limited to the distal half. seed and pollen strobili at all stages of Ovulate strobili cylindrical, typically solitary maturity. Further research is required in order but up to 3 per crown, erect at maturity, to determine whether there is any periodicity 8–15.5×4–4.5cm,sterileapex0.5–4cmand in the timing of the reproductive events of acute to acuminate, strobilus axis villous with this species. mixed rust-brown and white hairs, peduncles Ecology.—Larvae of a species of Eumaeus 15–50×1.2–2.7 cm, emerging white to cream Hübner were observed feeding on emergent tomentose, progressing maroon to brown Zamia leaves and live butterflies were tomentose, at maturity appearing green due to observed laying eggs on new leaves and loss of tomentum, megasporophylls in 7–10 emergent pollen strobili. Pollen strobili at orthostichies of 5–11 sporophylls each, stalk pollen release stage were observed harboring 8–10 mm long, villous with mixed rust-brown numerous clavicorn beetles of the genus 84 BRITTONIA [VOL 62 Pharaxonotha Reitt., known pollinators of two species have margins that are distinctly other Zamia species (Vovides, 1991; Tang, toothed on the upper half. The petiole and 1987; Stevenson et al., 1998). rachis of Z. pyrophylla is densely covered with Etymology.—The specific epithet refers to a ferruginous pubescence on young leaves and the glowing orange and red emergent leaves, have branched prickles, whereas those of Z. which progressively turn green from the apex cunaria and Z. ipetiensis are lightly covered to the base, giving them a fiery appearance. with a white pubescence and have unbranched Conservation status.—This species has prickles. The microsporophylls of Z. cunaria been collected only in an area covering and Z. ipetiensis are a uniform tan or cream approximately 25 km2 in the Municipalities color at pollen shed, whereas those of Z.